# Beyond HY5: COP1 Posttranslational Control of Anthocyanin Biosynthesis Proteins in Horticultural Crops

**Authors:** Gabriel Lasmar dos Reis, Agustín Zsögön, Antonio Chalfun-Junior, Lázaro Eustáquio Pereira Peres, Vagner Augusto Benedito

PMC · DOI: 10.3390/plants15040616 · Plants · 2026-02-14

## TL;DR

This paper explores how COP1, a protein involved in light signaling, controls anthocyanin production in plants beyond its known target HY5, with implications for improving crop nutrition.

## Contribution

The paper highlights novel COP1 targets and regulatory mechanisms in anthocyanin biosynthesis beyond HY5 in horticultural crops.

## Key findings

- COP1 regulates anthocyanin accumulation through multiple transcription factors and modules beyond HY5.
- Post-translational mechanisms involving COP1 are conserved across horticultural crops like apple and tomato.
- Understanding COP1's role could lead to improved nutritional traits in crops.

## Abstract

Anthocyanins are widespread specialized metabolites that provide pigmentation and antioxidant capacity, contributing to pollinator and seed-disperser attraction and to plant resistance to diverse environmental stresses. In human diets, anthocyanins are valued for their antioxidant and health-promoting properties. The biosynthetic pathway of anthocyanins is relatively conserved across plant species and is controlled by structural genes that encode the enzymes of the pathway along with regulatory genes, particularly transcription factors. This network integrates developmental and environmental signals, with light serving as a dominant cue: anthocyanins typically accumulate in light-exposed tissues and are repressed in darkness. A key node in this light-dependent switch is CONSTITUTIVE PHOTOMORPHOGENIC 1 (COP1), an E3 ubiquitin ligase that, in the dark, promotes polyubiquitination and proteasome-mediated turnover of positive regulators of anthocyanin production. Although ELONGATED HYPOCOTYL 5 (HY5) is a canonical COP1 target and major activator of anthocyanin biosynthesis, COP1 control of this pathway extends well beyond HY5. Evidence from Arabidopsis and multiple horticultural crops, including apple, pear, eggplant, and tomato, indicates that COP1 also regulates anthocyanin accumulation through interactions with additional transcription factors and regulatory modules. Here, we synthesize recent advances in COP1-centered regulation of anthocyanin biosynthesis, with an emphasis on post-translational mechanisms and COP1 targets beyond HY5. We also discuss emerging opportunities to leverage this regulatory axis for nutritional improvement in horticultural species.

## Linked entities

- **Genes:** COP1 (COP1 E3 ubiquitin ligase) [NCBI Gene 64326], RNY5 (RNA, Ro60-associated Y5) [NCBI Gene 6090]
- **Species:** Arabidopsis (taxon 3701)

## Full-text entities

- **Genes:** GSTF10 (glutathione S-transferase PHI 10) [NCBI Gene 817637] {aka ARABIDOPSIS THALIANA GLUTATHIONE S-TRANSFERASE PHI 10, ATGSTF10, ATGSTF4, EARLY DEHYDRATION-INDUCED 13, ERD13, F7F1.8}, 4CL2 (4-coumarate:CoA ligase 2) [NCBI Gene 821678] {aka 4-COUMARATE:COA LIGASE, 4-coumarate:CoA ligase 2, AT4CL2}, CIP7 (COP1-interacting protein 7) [NCBI Gene 828851] {aka COP1-interacting protein 7, F27G19.30, F27G19_30}, TT7 (Cytochrome P450 superfamily protein) [NCBI Gene 830693] {aka CYP75B1, CYTOCHROME P450 75B1, D501, F13G24.190, F13G24_190, F3'H}, DFR (dihydroflavonol 4-reductase) [NCBI Gene 544150] {aka FNR}, CO (B-box type zinc finger protein with CCT domain-containing protein) [NCBI Gene 831441] {aka B-box domain protein 1, BBX1, CONSTANS, F14F8.220, F14F8_220, FG}, FUS9 (Ubiquitin-conjugating enzyme family protein) [NCBI Gene 820557] {aka CIN4, CONSTITUTIVE PHOTOMORPHOGENIC 10, COP10, CYTOKININ-INSENSITIVE 4, E2 UBIQUITIN-CONJUGATING ENZYME, EMB144}, CRY1 (cryptochrome 1) [NCBI Gene 826470] {aka ATCRY1, BLU1, BLUE LIGHT UNINHIBITED 1, CRYPTOCHROME 1 APOPROTEIN (BLUE LIGHT PHOTORECEPTOR, ELONGATED HYPOCOTYL 4, HY4}, CHI (chitinase) [NCBI Gene 818959] {aka ''chitinase, F18O19.32, chitinase, putative, putative''}, COP1 (COP1 E3 ubiquitin ligase) [NCBI Gene 64326] {aka CFAP78, FAP78, RFWD2, RNF200}, TT4 (Chalcone and stilbene synthase family protein) [NCBI Gene 831241] {aka ATCHS, CHALCONE SYNTHASE, CHALCONE/STILBENE SYNTHASE, CHS, MAC12.28, MAC12_28}, SKP1 (S phase kinase-associated protein 1) [NCBI Gene 843928] {aka ARABIDOPSIS SKP1 HOMOLOGUE 1, ASK1, ATSKP1, S phase kinase-associated protein 1, SKP1A, T4O12.17}, MYB113 (myb domain protein 113) [NCBI Gene 842955] {aka AtMYB113, T27F4.12, T27F4_12, myb domain protein 113}, CRY2 (cryptochrome 2) [NCBI Gene 839529] {aka AT-PHH1, ATCRY2, CRYPTOCHROME 2 APOPROTEIN, F19P19.14, F19P19_14, FHA}, PAP1 (phosphatidic acid phosphatase 1) [NCBI Gene 814646] {aka ATLPP1, ATPAP1, F10A8.6, F10A8_6, LIPID PHOSPHATE PHOSPHATASE 1, LPP1}, PAP2 (Purple acid phosphatases superfamily protein) [NCBI Gene 837949] {aka F16A14.11, F16A14_11, purple acid phosphatase 2}, MUL1 (mitochondrial E3 ubiquitin protein ligase 1) [NCBI Gene 79594] {aka C1orf166, GIDE, MAPL, MULAN, RNF218}, FLS1 (flavonol synthase 1) [NCBI Gene 830765] {aka ATFLS1, FLAVONOL SYNTHASE, FLS, T2K12.5, flavonol synthase 1}, AT3G06710 (E3 ubiquitin ligase) [NCBI Gene 819856] {aka T8E24.2}, C4H (cinnamate-4-hydroxylase) [NCBI Gene 817599] {aka ATC4H, CINNAMATE 4-HYDROXYLASE, CINNAMATE-4-HYDROXYLASE, CYP73A5, REDUCED EPRDERMAL FLUORESCENCE 3, REF3}, F3H (flavanone 3-hydroxylase) [NCBI Gene 824287] {aka F3'H, FLAVANONE 3-HYDROXYLASE, TRANSPARENT TESTA 6, TT6, flavanone 3-hydroxylase}, DFR (dihydroflavonol 4-reductase) [NCBI Gene 834291] {aka DIHYDROFLAVONOL 4-REDUCTASE, DIHYDROKAEMPFEROL 4-REDUCTASE, M318, MJB21.18, MJB21_18, TT3}, UBC8 (ubiquitin conjugating enzyme 8) [NCBI Gene 834173] {aka ARABIDOPSIS THALIANA UBIQUITIN CONJUGATING ENZYME 8, ATUBC8, MBK23.24, MBK23_24, UBIQUITIN CONJUGATING ENZYME, ubiquitin conjugating enzyme 8}, HFR1 (basic helix-loop-helix (bHLH) DNA-binding superfamily protein) [NCBI Gene 839300] {aka FBI1, LONG HYPOCOTYL IN FAR-RED, REDUCED PHYTOCHROME SIGNALING 1, REDUCED SENSITIVITY TO FAR-RED LIGHT 1, REP1, RSF1}, COP1 (Transducin/WD40 repeat-like superfamily protein) [NCBI Gene 817857] {aka ARABIDOPSIS THALIANA CONSTITUTIVE PHOTOMORPHOGENIC 1, ATCOP1, CONSTITUTIVE PHOTOMORPHOGENIC 1, DEETIOLATED MUTANT 340, DET340, EMB168}, MYB90 (myb domain protein 90) [NCBI Gene 842957] {aka ATMYB90, PAP2, PRODUCTION OF ANTHOCYANIN PIGMENT 2, T27F4.14, T27F4_14, myb domain protein 90}, GL3 (basic helix-loop-helix (bHLH) DNA-binding superfamily protein) [NCBI Gene 834133] {aka GLABRA 3, GLABROUS 3, MYC6.2}, EGL3 (basic helix-loop-helix (bHLH) DNA-binding superfamily protein) [NCBI Gene 842669] {aka ATMYC-2, AtEGL3, EGL1, ENHANCER OF GLABRA 3, F24D7.16, F24D7_16}, PAP1 (production of anthocyanin pigment 1) [NCBI Gene 842120] {aka ATMYB75, F25P12.92, F25P12_92, MYB DOMAIN PROTEIN 75, MYB75, MYELOBLASTOSIS PROTEIN 75}, TT5 (Chalcone-flavanone isomerase family protein) [NCBI Gene 824678] {aka A11, AtCHI, CFI, CHALCONE FLAVANONE ISOMERASE, CHI, TRANSPARENT TESTA 5}, UF3GT (UDP-glucose:flavonoid 3-o-glucosyltransferase) [NCBI Gene 835489] {aka MJP23.2, MJP23_2, UDP-glucose:flavonoid 3-o-glucosyltransferase, UDP-glucosyl transferase 79B1, UGT79B1}, HY5 (Basic-leucine zipper (bZIP) transcription factor family protein) [NCBI Gene 830996] {aka BZIP TRANSCRIPTION FACTOR HY5, ELONGATED HYPOCOTYL 5, F2I11.150, F2I11_150, REVERSAL OF THE DET PHENOTYPE 5, TED 5}, TT8 (basic helix-loop-helix (bHLH) DNA-binding superfamily protein) [NCBI Gene 826571] {aka AtTT8, BHLH42, F17A8.170, F17A8_170, TRANSPARENT TESTA 8}, TTG1 (Transducin/WD40 repeat-like superfamily protein) [NCBI Gene 832523] {aka ATTTG1, K18P6.4, K18P6_4, TRANSPARENT TESTA GLABRA 1, TTG, UNARMED 23}, MYB [NCBI Gene 544113], PHYA (phytochrome A) [NCBI Gene 837483] {aka ELONGATED HYPOCOTYL 8, F14J9.23, F14J9_23, FAR RED ELONGATED 1, FAR RED ELONGATED HYPOCOTYL 2, FHY2}, COP8 (Proteasome component (PCI) domain protein) [NCBI Gene 834312] {aka ATS4, CONSTITUTIVE PHOTOMORPHOGENIC 14, CONSTITUTIVE PHOTOMORPHOGENIC 8, COP14, COP9 SIGNALOSOME SUBUNIT 4, CSN4}, PHYB (phytochrome B) [NCBI Gene 816394] {aka HY3, MSF3.17, MSF3_17, OOP1, OUT OF PHASE 1, PHYTOCHROME B}, LDOX (leucoanthocyanidin dioxygenase) [NCBI Gene 828387] {aka ANS, ANTHOCYANIDIN SYNTHASE, F7H19.60, F7H19_60, LEUCOANTHOCYANIDIN DIOXYGENASE, TANNIN DEFICIENT SEED 4}
- **Diseases:** injury to (MESH:D014947), inflammatory (MESH:D007249), diabetes (MESH:D003920), cancer (MESH:D009369)
- **Chemicals:** phosphorus (MESH:D010758), pelargonidin (MESH:C066957), metal (MESH:D008670), malonyl-CoA (MESH:D008316), cinnamic acid (MESH:C029010), MG132 (MESH:C072553), free radical (MESH:D005609), water (MESH:D014867), cyanidin 3-O-rutinoside (MESH:C428983), flavonol (MESH:C041477), malvidin (MESH:C065861), Dihydroflavonols (-), naringenin chalcone (MESH:C027329), peonidin (MESH:C473205), phenylalanine (MESH:D010649), cyanidin (MESH:C017154), delphinidin (MESH:C017185), naringenin (MESH:C005273), ATP (MESH:D000255), ROS (MESH:D017382), heavy metal (MESH:D019216), cyanidin 3-O-glucoside (MESH:C462279), flavonoid (MESH:D005419), Anthocyanin (MESH:D000872), petunidin (MESH:C473206), 4-coumaroyl-CoA. (MESH:C058644)
- **Species:** Prunus persica (peach, species) [taxon 3760], Malus domestica (apple, species) [taxon 3750], Pyrus pyrifolia (Asian pear, species) [taxon 3767], Pyrus communis (pear, species) [taxon 23211], Lactuca sativa (cultivated lettuce, species) [taxon 4236], Arabidopsis thaliana (mouse-ear cress, species) [taxon 3702], Solanum melongena (aubergine, species) [taxon 4111], Solanum lycopersicum (tomato, species) [taxon 4081], Homo sapiens (human, species) [taxon 9606]

## Full text

_Full body text omitted from this summary view._ Fetch the complete paper as Markdown: https://tomesphere.com/paper/PMC12944699/full.md

## Figures

3 figures with captions in the complete paper: https://tomesphere.com/paper/PMC12944699/full.md

## References

85 references — full list in the complete paper: https://tomesphere.com/paper/PMC12944699/full.md

---
Source: https://tomesphere.com/paper/PMC12944699